Manteodea Burmeister, 1829
Mantises are an order (Mantodea) of insects that contains over 2,400
species in about 430 genera in 15 families. The largest family is the
Mantidae ("mantids"). Mantises are distributed worldwide in temperate
and tropical habitats. They have triangular heads with bulging eyes
supported on flexible necks. Their elongated bodies may or may not
have wings, but all Mantodea have forelegs that are greatly enlarged
and adapted for catching and gripping prey; their upright posture,
while remaining stationary with forearms folded, has led to the common
name praying mantis.
The closest relatives of mantises are the termites and cockroaches
(Blattodea), which are all within the superorder Dictyoptera. Mantises
are sometimes confused with stick insects (Phasmatodea), other
elongated insects such as grasshoppers (Orthoptera), or other
unrelated insects with raptorial forelegs such as mantisflies
(Mantispidae). Mantises are mostly ambush predators, but a few
ground-dwelling species are found actively pursuing their prey. They
normally live for about a year. In cooler climates, the adults lay
eggs in autumn, then die. The eggs are protected by their hard
capsules and hatch in the spring. Females sometimes practice sexual
cannibalism, eating their mates after copulation.
Mantises were considered to have supernatural powers by early
civilizations, including Ancient Greece, Ancient Egypt, and Assyria. A
cultural trope popular in cartoons imagines the female mantis as a
femme fatale. Mantises are among the insects most commonly kept as
1 Taxonomy and evolution
2.3 Diet and predation
2.4 Antipredator adaptations
2.5 Reproduction and life history
2.6 Sexual cannibalism
3 Relationship with humans
3.1 In literature and art
3.2 Martial arts
3.3 In mythology and religion
3.4 As pets
3.5 For pest control
3.6 Mantis-like robot
5 External links
Taxonomy and evolution
See also: List of mantis genera and species
Over 2,400 species of mantis in about 430 genera are recognized.
They are predominantly found in tropical regions, but some live in
temperate areas. The systematics of mantises have long been
disputed. Mantises, along with stick insects (Phasmatodea), were once
placed in the order
Orthoptera with the cockroaches (now Blattodea)
and rock crawlers (now Grylloblattodea). Kristensen (1991) combined
the Mantodea with the cockroaches and termites into the order
Dictyoptera, suborder Mantodea. The name mantodea is formed from
the Ancient Greek words μάντις (mantis) meaning "prophet", and
εἶδος (eidos) meaning "form" or "type". It was coined in 1838 by
the German entomologist Hermann Burmeister. The order is
occasionally called the mantes, using a Latinized plural of Greek
mantis. The name mantid properly refers only to members of the family
Mantidae, which was, historically, the only family in the order. The
other common name, praying mantis, applied to any species in the
order, (though in Europe mainly to
Mantis religiosa), comes from
the typical "prayer-like" posture with folded fore limbs. The
vernacular plural "mantises" (used in this article) was confined
largely to the USA, with "mantids" predominantly used as the plural in
the UK and elsewhere, until the family
Mantidae was further split in
One of the earliest classifications splitting an all-inclusive
Mantidae into multiple families was that proposed by Beier in 1968,
recognizing eight families, though it was not until Ehrmann's
reclassification into 15 families in 2002 that a multiple-family
classification became universally adopted. Klass, in 1997, studied the
external male genitalia and postulated that the families Chaeteessidae
Metallyticidae diverged from the other families at an early
date. However, the
Thespidae are still both
considered polyphyletic, so the Mantodea will have to be revised.
Mantidflies (Mantispidae, related to lacewings, but unrelated to
mantises), have raptorial forelegs somewhat like those of mantises,
but their wings and antennae differ visibly.
The earliest mantis fossils are about 135 million years old, from
Siberia. Fossils of the group are rare: by 2007, only about 25
fossil species were known. Fossil mantises, including one from
Japan with spines on the front legs as in modern mantises, have been
Cretaceous amber. Most fossils in amber are nymphs;
compression fossils (in rock) include adults. Fossil mantises from the
Crato Formation in Brazil include the 10 mm (0.39 in) long
Santanmantis axelrodi, described in 2003; as in modern mantises, the
front legs were adapted for catching prey. Well-preserved specimens
yield details as small as 5 μm through X-ray computed tomography.
Because of the superficially similar raptorial forelegs, mantidflies
may be confused with mantises, though they are unrelated. Their
similarity is an example of convergent evolution; mantidflies do not
have tegmina (leathery forewings) like mantises, their antennae are
shorter and less thread-like, and the raptorial tibia is more muscular
than that of a similar-sized mantis and bends back further in
preparation for shooting out to grasp prey.
Wing arrangement of a typical mantis, adult male Raptrix perspicua
Mantises have large, triangular heads with a beak-like snout and
mandibles. They have two bulbous compound eyes, three small simple
eyes, and a pair of antennae. The articulation of the neck is also
remarkably flexible; some species of mantis can rotate their heads
nearly 180°. The mantis thorax consists of a prothorax, a
mesothorax, and a metathorax. In all species apart from the genus
Mantoida, the prothorax, which bears the head and forelegs, is much
longer than the other two thoracic segments. The prothorax is also
flexibly articulated, allowing for a wide range of movements of the
head and fore limbs while the remainder of the body remains more or
The raptorial foreleg, showing the unusually long coxa, which,
together with the trochanter, gives the impression of a femur. The
femur itself is the proximal segment of the grasping part of the leg.
Mantises have two spiked, grasping forelegs ("raptorial legs") in
which prey items are caught and held securely. In most insect legs,
including the posterior four legs of a mantis, the coxa and trochanter
combine as an inconspicuous base of the leg; in the raptorial legs,
however, the coxa and trochanter combine to form a segment about as
long as the femur, which is a spiky part of the grasping apparatus
(see illustration). Located at the base of the femur is a set of
discoidal spines, usually four in number, but ranging from none to as
many as five depending on the species. These spines are preceded by a
number of tooth-like tubercles, which, along with a similar series of
tubercles along the tibia and the apical claw near its tip, give the
foreleg of the mantis its grasp on its prey. The foreleg ends in a
delicate tarsus used as a walking appendage, made of four or five
segments and ending in a two-toed claw with no arolium.
Mantises can be loosely categorized as being macropterous
(long-winged), brachypterous (short-winged), micropterous
(vestigial-winged), or apterous (wingless). If not wingless, a mantis
has two sets of wings: the outer wings, or tegmina, are usually narrow
and leathery. They function as camouflage and as a shield for the hind
wings, which are clearer and more delicate. The abdomen of all
mantises consists of 10 tergites, with a corresponding set of nine
sternites visible in males and seven visible in females. The abdomen
tends to be slimmer in males than females, but ends in a pair of cerci
in both sexes.
Head of Archimantis latistyla, showing the compound eyes and labrum
Mantises have stereo vision. They locate their prey by
sight; their compound eyes contain up to 10,000 ommatidia. A small
area at the front called the fovea has greater visual acuity than the
rest of the eye, and can produce the high resolution necessary to
examine potential prey. The peripheral ommatidia are concerned with
perceiving motion; when a moving object is noticed, the head is
rapidly rotated to bring the object into the visual field of the
fovea. Further motions of the prey are then tracked by movements of
the mantis's head so as to keep the image centered on the
fovea. The eyes are widely spaced and laterally situated,
affording a wide binocular field of vision and precise stereoscopic
vision at close range. The dark spot on each eye that moves as it
rotates its head is a pseudopupil. This occurs because the ommatidia
that are viewed "head-on" absorb the incident light, while those to
the side reflect it.
As their hunting relies heavily on vision, mantises are primarily
diurnal. Many species, however, fly at night, and then may be
attracted to artificial lights. Mantises in the family Liturgusidae
collected at night have been shown to be predominately males; this
is probably true for most mantises. Nocturnal flight is especially
important to males in locating less-mobile females by detecting their
pheromones. Flying at night exposes mantises to fewer bird predators
than diurnal flight would. Many mantises also have an auditory
thoracic organ that helps them avoid bats by detecting their
echolocation calls and responding evasively.
Diet and predation
Mantis eating a common bushbrown butterfly, Mycalesis perseus
Mantises are generalist predators of arthropods. The majority of
mantises are ambush predators that only feed upon live prey within
their reach. They either camouflage themselves and remain stationary,
waiting for prey to approach, or stalk their prey with slow, stealthy
movements. Larger mantises sometimes eat smaller individuals of
their own species, as well as small vertebrates such as lizards,
frogs, and small birds.
Tenodera sinensis feeding on a cricket
Most mantises stalk tempting prey if it strays close enough, and will
go further when they are especially hungry. Once within reach,
mantises strike rapidly to grasp the prey with their spiked raptorial
forelegs. Some ground and bark species pursue their prey in a more
active way. For example, members of a few genera such as the ground
mantises, Entella, Ligaria, and Ligariella run over dry ground seeking
prey, much as tiger beetles do.
Ligariella, a ground mantid that runs down its prey
The fore gut of some species extends the whole length of the insect
and can be used to store prey for digestion later. This may be
advantageous in an insect that feeds intermittently. Chinese
mantises live longer, grow faster, and produce more young when they
are able to eat pollen.
Further information: flower mantis
Mantises are preyed on by vertebrates such as frogs, lizards, and
birds, and by invertebrates such as spiders, large species of hornets,
and ants. Some hunting wasps, such as some species of Tachytes
also paralyse some species of mantis to feed their young.
Generally, mantises protect themselves by camouflage, most species
being cryptically colored to resemble foliage or other backgrounds,
both to avoid predators and to better snare their prey. Those that
live on uniformly colored surfaces such as bare earth or tree bark are
dorsoventrally flattened so as to eliminate shadows that might reveal
their presence. The species from different families called flower
mantises are aggressive mimics: they resemble flowers convincingly
enough to attract prey that come to collect pollen and
nectar. Some species in Africa and Australia are able to
turn black after a molt towards the end of the dry season; at this
time of year, bush fires occur and this coloration enables them to
blend in with the fire-ravaged landscape (fire melanism).
Aggressive mimicry: Malaysian orchid mantises are camouflaged pink or
yellow, matching the coloration of local orchids.
When directly threatened, many mantis species stand tall and spread
their forelegs, with their wings fanning out wide. The fanning of the
wings makes the mantis seem larger and more threatening, with some
species enhancing this effect with bright colors and patterns on their
hind wings and inner surfaces of their front legs. If harassment
persists, a mantis may strike with its forelegs and attempt to pinch
or bite. As part of the bluffing (deimatic) threat display, some
species may also produce a hissing sound by expelling air from the
abdominal spiracles. Mantises lack chemical protection, so their
displays are largely bluff. When flying at night, at least some
mantises are able to detect the echolocation sounds produced by bats;
when the frequency begins to increase rapidly, indicating an
approaching bat, they stop flying horizontally and begin a descending
spiral toward the safety of the ground, often preceded by an aerial
loop or spin. If caught, they may slash captors with their raptorial
Mantises, like stick insects, show rocking behavior in which the
insect makes rhythmic, repetitive side-to-side movements. Functions
proposed for this behavior include the enhancement of crypsis by means
of the resemblance to vegetation moving in the wind. However, the
repetitive swaying movements may be most important in allowing the
insects to discriminate objects from the background by their relative
movement, a visual mechanism typical of animals with simpler sight
systems. Rocking movements by these generally sedentary insects may
replace flying or running as a source of relative motion of objects in
the visual field. As ants may be predators of mantises, genera
such as Loxomantis, Orthodera, and Statilia, like many other
arthropods, avoid attacking them. Exploiting this behavior, a variety
of arthropods, including some early-instar mantises, mimic ants to
evade their predators.
Choeradodis has leaf-like fore wings and a widened green
Iris oratoria performs a bluffing threat display, rearing
back with the forelegs and wings spread and mouth opened.
The jeweled flower mantis, Creobroter gemmatus: the brightly colored
wings are opened suddenly in a deimatic display to startle predators.
Some mantis nymphs mimic ants to avoid predators.
Reproduction and life history
The mating season in temperate climates typically takes place in
autumn, while in tropical areas, mating can occur at any time
of the year. To mate following courtship, the male usually leaps
onto the female's back, clasping her thorax and wing bases with his
forelegs. He then arches his abdomen to deposit and store sperm in a
special chamber near the tip of the female's abdomen. The female lays
between 10 and 400 eggs, depending on the species. Eggs are typically
deposited in a froth mass-produced by glands in the abdomen. This
froth hardens, creating a protective capsule, which together with the
egg mass is called an ootheca. Depending on the species, the ootheca
can be attached to a flat surface, wrapped around a plant, or even
deposited in the ground. Despite the versatility and durability of the
eggs, they are often preyed on, especially by several species of
parasitoid wasps. In a few species, mostly ground and bark mantises in
the family Tarachodidae, the mother guards the eggs. The cryptic
Tarachodes maurus positions herself on bark with her abdomen covering
her egg capsule, ambushing passing prey and moving very little until
the eggs hatch. An unusual reproductive strategy is adopted by
Brunner's stick mantis from the southern United States; no males have
ever been found in this species, and the females breed
parthenogenetically. The ability to reproduce by parthenogenesis
has been recorded in at least two other species, Sphodromantis viridis
and Miomantis sp., although these species usually reproduce
sexually. In temperate climates, adults do not survive the
winter and the eggs undergo a diapause, hatching in the spring.
As in closely related insect groups in the superorder Dictyoptera,
mantises go through three life stages: egg, nymph, and adult (mantises
are among the hemimetabolous insects). For smaller species, the eggs
may hatch in 3–4 weeks as opposed to 4–6 weeks for larger species.
The nymphs may be colored differently from the adult, and the early
stages are often mimics of ants. A mantis nymph grows bigger as it
molts its exoskeleton. Molting can happen five to 10 times before the
adult stage is reached, depending on the species. After the final
molt, most species have wings, though some species remain wingless or
brachypterous ("short-winged"), particularly in the female sex. The
lifespan of a mantis depends on the species; smaller ones may live
4–8 weeks, while larger species may live 4–6 months.
Mantis religiosa mating (brown male, green female)
Stagmomantis carolina laying ootheca
Recently laid ootheca by
Newly hatched mantises
Further information: Sexual cannibalism
Sexual cannibalism in
Sexual cannibalism is common among most predatory species of mantises
in captivity. It has sometimes been observed in natural populations,
where about a quarter of male-female encounters result in the male
being eaten by the female. Around 90% of the predatory
species of mantises participate in sexual cannibalism. Adult males
typically outnumber females at first, but their numbers may be fairly
equivalent later in the adult stage, possibly because females
selectively eat the smaller males. In Tenodera sinensis, 83% of
males escape cannibalism after an encounter with a female, but since
multiple matings occur, the probability of a male's being eaten
The female may begin feeding by biting off the male's head (as they do
with regular prey), and if mating has begun, the male's movements may
become even more vigorous in its delivery of sperm. Early researchers
thought that because copulatory movement is controlled by a ganglion
in the abdomen, not the head, removal of the male's head was a
reproductive strategy by females to enhance fertilization while
obtaining sustenance. Later, this behavior appeared to be an artifact
of intrusive laboratory observation. Whether the behavior is natural
in the field or also the result of distractions caused by the human
observer remains controversial. Mantises are highly visual organisms
and notice any disturbance in the laboratory or field, such as bright
lights or moving scientists. Chinese mantises that had been fed ad
libitum (so that they were not hungry) actually displayed elaborate
courtship behavior when left undisturbed. The male engages the female
in a courtship dance, to change her interest from feeding to
mating. Under such circumstances, the female has been known to
respond with a defensive deimatic display by flashing the colored
eyespots on the inside of her front legs.
The reason for sexual cannibalism has been debated; experiments show
that females on low quality diets have a higher chance to engage in
sexual cannibalism compared to females on high quality diets. Some
consider that submissive males gain a selective advantage by producing
offspring. This theory is supported by a quantifiable increase in the
duration of copulation among males which are cannibalized, in some
cases doubling both the duration and the chance of fertilization. This
is contrasted by a study where males were seen to approach hungry
females with more caution, and were shown to remain mounted on hungry
females for a longer time, indicating that males that actively avoid
cannibalism may mate with multiple females. The same study also found
that hungry females generally attracted fewer males than those that
were well fed. The act of dismounting after copulation is
dangerous for males, for at this time, females most frequently
cannibalize their mates. An increase in mounting duration appears to
indicate that males wait for an opportune time to dismount a hungry
female, who would be likely to cannibalize her mate.
Relationship with humans
In literature and art
One of the earliest mantis references is in the ancient Chinese
dictionary Erya, which gives its attributes in poetry, where it
represents courage and fearlessness, and a brief description. A later
text, the Jingshi Zhenglei Daguan Bencao ("Great History of Medical
Material Annotated and Arranged by Types, Based upon the Classics and
Historical Works") from 1108, gives accurate details of the
construction of the egg packages, the development cycle, anatomy, and
the function of the antennae. Although mantises are rarely mentioned
in Ancient Greek sources, a female mantis in threat posture is
accurately illustrated on a series of fifth-century BC silver coins,
including didrachms, from
Metapontum in Sicily. In the 10th
Byzantine era Adages, Suidas describes an insect
resembling a slow-moving green locust with long front legs. He
Zenobius 2.94 with the words seriphos (maybe a mantis) and
graus, an old woman, implying a thin, dried-up stick of a body.
Western descriptions of the biology and morphology of the mantises
became more accurate in the 18th century. Roesel von Rosenhof
illustrated and described mantises and their cannibalistic behavior in
the Insekten-Belustigungen (
Aldous Huxley reflected on death as a pair of Gongylus
Aldous Huxley made philosophical observations about the nature of
death while two mantises mated in the sight of two characters in his
1962 novel Island (the species was Gongylus gongylodes). The
naturalist Gerald Durrell's humorously autobiographical 1956 book My
Family and Other Animals includes a four-page account of an almost
evenly matched battle between a mantis and a gecko. Shortly before the
fatal dénouement, Durrell narrates:
he [Geronimo the gecko] crashed into the mantis and made her reel, and
grabbed the underside of her thorax in his jaws. Cicely [the mantis]
retaliated by snapping both her front legs shut on Geronimo's
hindlegs. They rustled and staggered across the ceiling and down the
wall, each seeking to gain some advantage.
M. C. Escher's woodcut Dream depicts a human-sized mantis standing on
a sleeping bishop. The 1957 film
The Deadly Mantis
The Deadly Mantis features a
mantis as a giant monster.
A cultural trope imagines the female mantis as a femme fatale. The
idea is propagated in cartoons by Cable, Guy and Rodd, LeLievre, T.
McCracken, and Mark Parisi, among others. It ends
Isabella Rossellini's short film about the life of a praying mantis in
Green Porno season for the Sundance Channel.
Grandmasters of the Shaolin Temple, Shi DeRu and Shi DeYang,
Southern Praying Mantis
Southern Praying Mantis style of martial art
Two martial arts separately developed in China have movements and
fighting strategies based on those of the mantis. As one of
these arts was developed in northern China, and the other in southern
parts of the country, the arts are nowadays referred to (both in
English and Chinese) as 'Northern Praying Mantis' and 'Southern
Praying Mantis'. Both are very popular in China, and have also
been exported to the West in recent decades.
In mythology and religion
The mantis was revered by the southern African
Khoi and San in whose
cultures man and nature were intertwined; for its praying posture, the
mantis was even named Hottentotsgot ("god of the Khoi") in the
Afrikaans language that had developed among the first European
settlers. However, at least for the San, the mantis was only one
of the manifestations of a trickster-deity who could assume many other
forms, such as a snake, hare or vulture. Several ancient
civilizations did consider the insect to have supernatural powers; for
the Greeks, it had the ability to show lost travelers the way home; in
the Ancient Egyptian
Book of the Dead
Book of the Dead the "bird-fly" is a minor god
that leads the souls of the dead to the underworld; in a list of
Nineveh grasshoppers (buru), the mantis is named
necromancer (buru-enmeli) and soothsayer (buru-enmeli-ashaga).
Gray adult female
Carolina mantis in human hand
Mantises are among the insects most widely kept as pets.
Because the lifespan of a mantis is only about a year, people who want
to keep mantises often breed them. In 2013 at least 31 species were
kept and bred in the United Kingdom, the Netherlands, and the United
States. In 1996 at least 50 species were known to be kept in
captivity by members of the
Mantis Study Group. The Independent
described the "giant Asian praying mantis" as "part stick insect with
a touch of Buddhist monk", and stated that they needed a vivarium
around 30 cm (12 in) on each side. The Daily South
argued that a pet insect was no weirder than a pet rat or ferret, and
that while a pet mantis was unusual, it would not "bark, shed, [or]
need shots or a litter box".
For pest control
Gardeners who prefer to avoid pesticides may encourage mantises in the
hope of controlling insect pests. However, mantises do not have
key attributes of biological pest control agents; they do not
specialize in a single pest insect, and do not multiply rapidly in
response to an increase in such a prey species, but are general
predators. They eat whatever they can catch, including both
harmful and beneficial insects. They therefore have "negligible
value" in biological control.
Two species, the
Chinese mantis and the European mantis, were
deliberately introduced to North America in the hope that they would
serve as pest controls for agriculture; they have spread widely in
both the United States and Canada.
Further information: Biomimicry
A prototype robot inspired by the forelegs of the praying mantis has
front legs that allow the robot to walk, climb steps, and grasp
objects. The multi-jointed leg provides dexterity via a rotatable
joint. Future models may include a more spiked foreleg to improve the
grip and ability to support more weight.
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Mantis Study Group – Information on mantises, phylogenetics and
Archaeognatha (jumping bristletails)
Thysanura (Zygentoma) (silverfish, firebrats)
Odonata (dragonflies, damselflies)
Phasmatodea (stick and leaf insects)
Notoptera (ice-crawlers, gladiators)
Orthoptera (crickets, wetas, grasshoppers, locusts)
Zoraptera (angel insects)
Blattodea (cockroaches, termites)
Psocodea (barklice, lice)
Hemiptera (cicadas, aphids, true bugs)
Hymenoptera (sawflies, wasps, ants, bees)
Strepsiptera (twisted-winged parasites)
Megaloptera (alderflies, dobsonflies, fishflies)
Neuroptera (net-winged insects: lacewings, mantidflies, antlions)
Mecoptera (scorpionflies) + Siphonaptera (fleas)
Diptera (gnats, mosquitoes, flies)
Lepidoptera (moths, butterflies)
Four most speciose orders are marked in bold
Italic are paraphyletic groups
Based on Sasaki et al. (2013)
Extinct incertae sedis families and genera are marked in italic
Extant Mantodea families
Hymenopodidae (flower mantises)
List of mantis genera and species
Insects in culture
In the arts
Insects in art
Insects in film
Insects in literature
Insects in music
List of insect-inspired songs
Insects on stamps
Insects in religion
Colorado potato beetle
Cottony cushion scale
Western corn rootworm
Insect bites and stings
Insect sting allergy
House longhorn beetle
Home-stored product entomology
Alfred Russel Wallace
Hans Zinsser (Rats, Lice and History)
Lafcadio Hearn (
Living things in culture
Fauna Europaea: 11918